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Revisiting the Cambrian Burgess Shale Palaeocommunity in Light of New Field Discoveries from Marble Canyon, Kootenay National Park, British Columbia

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Revisiting the Cambrian Burgess Shale Palaeocommunity in Light of New Field Discoveries from Marble Canyon, Kootenay National Park, British Columbia Revisiting the Cambrian Burgess Shale Palaeocommunity in Light of New Field Discoveries from Marble Canyon, Kootenay National Park, British Columbia by Karma Nanglu A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy Department of Ecology and Evolutionary Biology University of Toronto © Copyright by Karma Nanglu 2019 Revisiting the Cambrian Burgess Shale palaeocommunity in light of new field discoveries from Marble Canyon, Kootenay National Park, British Columbia Karma Nanglu Doctor of Philosophy Department of Ecology and Evolutionary Biology University of Toronto 2019 ABSTRACT The 508-million-year-old Burgess Shale (British Columbia) is among the most important fossil localities in the world as it provides a direct window into the Cambrian Explosion, the phenomenon whereby most metazoan groups appeared rapidly in the fossil record for the first time. For over 100 years, Burgess Shale fossils have provided unique insights into the early evolutionary history of animal life, but a holistic description of the community ecology of the Burgess Shale has remained elusive. This dissertation aims to reinvestigate the Burgess Shale paleocommunity in light of the recently discovered Marble Canyon fossil site in Kootenay National Park, integrating this new dataset with those from the type areas in Yoho National Park, 40km to the northwest (Walcott Quarry, Raymond Quarry, and Tulip Beds). The first three chapters of this dissertation focus on the organismal level. An experimental decay study (Chapter 1) provides a framework for the interpretation of taphonomic bias in community reconstructions. The redescription of three enigmatic Cambrian taxa (Chapters 2 and 3) provide novel phylogenetic and ecological information for understanding the patterns of diversity and niche structure at Marble Canyon. The community analysis (Chapter 4) represents the largest quantitative study of the Burgess Shale to date. Patterns of faunal stasis between most adjacent bedding assemblages followed by periodic variations in abundance and ii species identity are found at both Marble Canyon and Walcott Quarry, the two best-studied sites with the finest level of stratigraphic/temporal data. Across the entire Burgess Shale paleocommunity, major shifts in representative taxonomic groups and ecological modes occur between major localities. The results of this work suggest that the Burgess Shale as a paleocommunity was highly heterogeneous in both its taxonomic and ecological composition. Shifting abiotic and environmental variables are determined to be more significant in the long- term structuring of Burgess Shale communities than species interactions such as predation, which was previously considered paramount. iii ACKNOWLEDGEMENTS The first thanks must go to my supervisor, Jean-Bernard Caron. To have unfettered access to what is, in my obviously unbiased opinion, one of the most significant and exciting fossil discoveries in the last half-century has been the opportunity of a lifetime, and I cannot over express my thanks to Jean- Bernard for trusting me with this thesis. Moreover, to spend three months in the Burgess Shale and to be part of the process of paleontology from prospecting to discovery to analysis to seeing my work in the galleries of the Royal Ontario Museum has been an experience that is impossible to compare. When I reflect on the core insight I’ve come away from this PhD with, the single word that comes to mind is scale. Unlike many of the other grad students in our department, being a paleontologist was not my childhood dream (forgiving, for a moment, the standard childhood shortsighted dalliance with dinosaurs), and so I was perhaps less prepared for thinking about deep time than most of my peers when I started. Working on the Cambrian explosion, then, is truly to be thrown in into the temporal deep end. I've come to realize that to work as a paleontologist is as close as you can come to being a time traveler, and I thank Jean-Bernard once again for giving me that chance. I would like to acknowledge my committee members Dr. Hélène Cyr and Dr. Doug Currie. Their feedback and support for my research over the last 5 years has been an essential component of becoming the researcher I am now. The Invertebrate Paleontology department at the Royal Ontario Museum has been a second home to me for the last 5 years. While the support of the entire museum has helped me get to where I am today, I’ll quickly address each of various invertebrate workers in turn that I’ve had the pleasure of knowing: Pete Fenton – Before I had even technically started as a graduate student, Jean-Bernard put me in contact with Pete as the be-all end-all problem fixer for my future studies. I put that reputation to the test in short order; I’ve since been told that our Paleobiology department rarely requires chambers for euthanizing, chilling, and rotting worms. This may explain our lack of a readily available incubator. Pete’s ability to iv channel Macgyver left me with a faith in his problem solving abilities that would go unshaken to this day; the experiments for the first chapter of my thesis were facilitated by a bucket of water, an electric fan and a bag of rocks. Perhaps more importantly than that, Pete has always been an unfailingly supportive person to work with, and often a great sympathetic ear when academia has set you down the path of insanity. Cédric Aria – I don’t think it would be a stretch to say that Cédric and I have had a disagreement or two in the past. But I’m certainly no stranger to arguments being my primary mode of communication and so in that way, I think we share a common language. I came to truly appreciate our conversations, and I can think of few people who put as much of themselves into their work as Cédric. There is a real sense of the auteur to Cédric, a wholehearted feeling that what he’s working on is of critical importance and that he’ll brook no compromise in what he thinks is the truth. I think that’s something that we can all admire, even if we don’t always agree. He’s also got a pretty good singing voice, and his mountainside yodeling is not to be despised. Joe Moysiuk - I’m still sore that I didn’t make it into the acknowledgments for Joe’s first paper. But that’s only because I want to lay some claim to his future successes, which I’m sure will be manifold. The guy is pretty much designed to work on the Cambrian, it’s uncanny. He’s also a solid confidante during field work, which as anyone who’s spent time in the Burgess Shale will tell you, is absolutely crucial. Sometimes, after a long day of back-breaking hiking, all you need is to be reminded of a burnt risotto to put your life back into perspective. Maryam Akrami – I’m not sure that I love fossils half as much as Maryam. Sometimes I don’t know if anyone loves fossils half as much as Maryam. There have been more than a few moments where, feeling beaten down by grad school, I’ve been stunned by how much raw appreciation Maryam has for the material we’re working on. It really re-centers the importance of this type of work, which you can sometimes lose sight of among all the meetings and deadlines. I’d also say our world views differ pretty v significantly, but I think Maryam is a person you can trust to speak her mind and stand by her convictions, which is something I really respect about her. I would also like to thank Sebastian Kvist. I’ve learned a lot from Sebastian, and teaching the invertebrate diversity labs with him for the last three years has been a huge joy and privilege. Hanging out with him and his lab at the ROM has felt like spending time with extended family, and I can think of few people who are as unfailingly supportive. It’s really great to have a friend who you can trust will always be in your corner, and Sebastian is that kind of guy. Grad school was the first time in my life where I felt like I was on the same wavelength as the people around me. I'd never had a group of peers who were as excited as I was for science, or wanted to spend their off-hours just reading about the most bizarre aspects of animal life. There are many to thank, but in alphabetical order I wanted to recognize: Erika Anderson, Sean Anderson, Victoria Arbour, Viviana Astudillo, James Boyko, Rowshyra Castañeda, Kentaro Chiba, Hollis Dahn, Danielle Dufault, Rafael Eiji Iwama, Michael Foisy, Bryan Gee, Scott Gilchrist, Jessica Hawthorn, Derek Larson, Aaron LeBlanc, Mark MacDougall, Claire Manglicmot, Melanie Massey, Mateus Pepinelli, Ashley Reynolds, Jade Simon, and Sarah Steele. Special thanks are also due to three fellow grad students in particular. Melissa Orobko was a great support for me during many of the most difficult moments during my PhD. I recall one particularly long and unpleasant walk through the snow while I very seriously questioned whether or not I had it in me to stick it out in grad school. Melissa helped talk me down from that metaphorical ledge. Thomas M. Cullen fits into a rare niche amongst the people I've met throughout grad school. That niche is being someone who I simultaneously respect as a peer, but also admire as an academic goal to strive towards. There are few people who I have been able to go to for such consistently good advice; you couldn’t ask for a better sounding board or friend.
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